Managed realignment is now widely seen as an important part of coastal management strategy, as an environmentally sustainable, cost-effective alternative to traditional, hard defences. However, the trajectory of salt marsh development in managed realignment schemes remains uncertain and it is unclear how sites should be managed to fulfil both coastal defence and biodiversity objectives. In this study, the overarching aim is to find out ways in which we can create salt marshes that are self-sustaining and function as closely to natural ecosystems as possible, by considering the linked biogeomorphological processes in salt marsh ecosystems. Such an approach will improve predictions of habitat development and recommendations for future practice in managed realignment schemes. The first important question is: How do pre-alignment plant species growing within managed realignment sites respond to salt water inundation upon re-exposure to flooding? Understanding the responses of the terrestrial vegetation community to initial seawater flooding may improve predictions of the short-term transition into salt marsh vegetation. To answer this, the effects of seawater inundation on pre-existing vegetation are initially examined in a greenhouse experiment in Chapter 3. It was demonstrated that one common plant species component of many coastal grasslands, Trifolium repens, responds poorly to simulated seawater soil flooding, but the response is population-, i.e. ecotype-, specific; therefore, the species consequently has an adaptive capacity to withstand short periods of soil inundation by seawater. In addition, I look at how and why the vegetation community of a restored site transitions following the reintroduction of tidal water, including the response of the original community of non-salt marsh plant species to salt water inundation and subsequent salt marsh plant community reassembly. After three years of tidal inundation at South Efford managed realignment site (SEM), terrestrial vegetation had decreased in cover and nearly all species recorded on the adjacent natural marsh had colonised. However, the cover of salt marsh species was limited by waterlogging, caused by modifications to the tidal regime by a self-regulating tidal gate. This leads on to the second question: How do new engineering techniques alter the tidal regime and what specific aspects of the new regime drive plant community reassembly and sedimentation patterns? In Chapters 4, 5 and 6, three years of ecological and geomorphological development are investigated in response to a variable inundation regime imposed by regulated tidal exchange at SEM. Inundation of the marsh surface was very regular, but water levels were not deep enough to encourage sufficient morphological development, sedimentation nor hydrochory. In contrast, ecological development was limited by waterlogging. Balancing the tidal regime with the drainage efficiency of managed realignment sites may be the most likely scenario under which restored salt marsh will develop with maximum biodiversity benefits. Otherwise, further management techniques, such as the excavation of tidal channels, may need to be employed to improve site drainage. Consequently, the final question is: How can biodiversity be maximized on realignment sites through the use of different management techniques and site design? Tidal channels on a range of managed and natural sites were shown to improve the drainage efficiency of adjacent soils (particularly channels of greater width and/or higher Strahler order). Plant species diversity was generally higher on channel banks in managed realignment sites. On sites with highly reduced soils, the colonisation and establishment of halophytes could be advanced on the banks of tidal channels. Additionally, topographic heterogeneity introduced by tidal channels created a variety of habitat niches, which allowed a range of salt marsh species to establish in the absence of highly competitive species, such as Elymus repens. Results from this study could contribute to the generation of a number of recommendations for the implementation of managed realignment schemes, particularly regarding the excavation of tidal creek networks.

There is a paucity of published information on fish utilization of salt marshes outside North America. This dissertation represents the first intensive examination of the ichthyofauna associated with salt marshes in southern Africa and examines the species composition, diversity, size structure, distribution and estuarine dependence of fishes that utilize salt marshes in the Kariega Estuary. The research was concentrated on Taylor's salt marsh, with comparative studies being conducted on other salt marshes and habitats within the marine dominated Kariega Estuary. The possible importance of salt marshes as feeding and refuge areas for fishes was examined, as was the role of salt marshes in the food web of the estuary. The fishes frequenting salt marshes in the Kariega Estuary were predominantly the juveniles of marine species, with Mugilidae being the dominant family. The ichthyofauna was distributed primarily in the intertidal creeks with very few specimens captured on the vegetated Spartina maritima and Sarcocomia perennis flats. The different reaches of the intertidal creek were characterised by distinct fish assemblages. The fish assemblages associated with the intertidal salt marsh creeks were significantly different from those found in the eelgrass beds, the other dominant intertidal habitat in the Kariega Estuary. The eelgrass beds were dominated by estuarine fish species and had a higher density and standing stock of fishes when compared to the salt marsh creeks. The diversity of fishes in the two habitats was however similar. The intertidal salt marsh creek ichthyofauna also differed significantly from that found in the main estuary channel. The creek ichthyofauna was dominated by 0+ juveniles while the main channel had many subadult and adult fishes. The channel habitat also had numerous large piscivorous fishes which were absent from the salt marsh creeks. The low number of piscivorous fishes, together with limited fish predation from other sources, may be the reason why salt marshes provide a refuge for juvenile fishes that frequent these habitats. Unlike previous studies on North American, Australian and European salt marshes, the dominant fish species that frequented Taylor's marsh were not recorded feeding extensively on the marsh, and those that did had a limited distribution. The role of fishes in the transfer of energy off the Kariega salt marshes is therefore likely to be minimal. A stable carbon isotope study on the dominant primary producers and consumers within the Kariega Estuary revealed that detritus originating from the high lying salt marsh plants Sarcocornia perennis and Chenolea diffusa was not utilized by fishes in the Kariega Estuary. Preliminary results indicated that the cord grass Spartina maritima may be an important energy source to the fishes in the Kariega Estuary, but further research is needed to confirm this.

A comprehensive field and laboratory study of nitrogen uptake in intertidal seaweeds was undertaken. Methods for measuring nitrogen uptake rates were evaluated. Short initial periods of rapid ammonium uptake were common in nitrogen deficient plants. The presence of ammonium inhibited nitrate uptake, but a certain degree of nitrogen starvation overcame this suppression. Laboratory studies with Porphyra perforata showed that nitrogen starved cultures maintained rapid initial ammonium uptake rates. The nitrate uptake system did not remain activated. Nitrogen starvation also resulted in a general decrease in soluble nitrogen content and a transient increase in nitrate reductase activity. The effectiveness of in vitro and in vivo nitrate reductase assays was investigated. The rate of nitrite production in the in vivo assay varied with incubation time. Therefore, the in vitro assay was used. Nitrate grown cultures of Porphyra perforata maintained high ammonium uptake rates. It was suggested that the rate of nitrate reduction was limiting the supply of nitrogen for further assimilation which may control ammonium uptake. Ammonium arid ammonium plus nitrate grown cultures had very low nitrogen uptake rates and nitrate reductase activities. Field studies with Gracilaria verrucosa confirmed that growth on ammonium inhibited nitrate uptake, nitrate accumulation and nitrate reductase activity. The presence of ammonium did not inhibit nitrate uptake rates in severely starved populations. All populations maintained high ammonium uptake rates suggesting that they were nitrogen limited at this time (August). Ammonium and nitrate uptake were saturable in the high intertidal G. verrucosa population but not in the low intertidal population. An investigation was made into the effect of nitrogen source and periodic exposure to air on growth, development and nitrogen uptake in Fucus distichus germlings. Gamete release, fertilization, germination and germling growth had no requirement for a specific form of nitrogen. Periodic exposure to air increased secondary rhizoid development. Ammonium and nitrate uptake rates of the germlings were much higher than for the mature thalli, but the affinity for nitrate was similar. The germlings showed saturable uptake kinetics but the mature thalli did not. The presence of ammonium inhibited nitrate uptake by the mature plants but not by the germlings. Mild desiccation enhanced nutrient uptake rates in several intertidal seaweeds. This uptake response occurred when growth was limited by that particular nutrient and when the thallus had been exposed to periodic desiccation for several weeks. The degree of enhancement, the percent desiccation producing maximum uptake rates and the tolerance to higher degrees of desiccation were related to intertidal location. This was shown to be an intraspecific as well as an interspecific adaptation. Transplant experiments with G. verrucosa showed that enhanced nutrient uptake rates after desiccation were related to intertidal height and not geographic location and that this response could be induced in approximately five weeks. It was suggested that this enhanced uptake response was an adaptation to nitrogen procurement and C/N homeostasis following periodic exposure when carbon was assimilated but when other nutrients were not available. / Science, Faculty of / Botany, Department of / Graduate

Coastal bays and salt marshes are buffer zones located at the interface between land and ocean, and provide ecologically and commercially important services worldwide. Unfortunately, their location makes them vulnerable and sensitive to sea-level rise (SLR), reduced sediment loads and anthropogenic modifications of the shoreline. Sediment budget and sediment availability are direct metrics for evaluating the resilience of salt marshes and coastal bays to various stressors (e.g. SLR). Salt marshes requires adequate sediment inputs to maintain their elevation with respect to sea level. Understanding sediment trajectories, sediment fluxes and sediment trapping capacities in different geomorphic unit facilitates efficient restorations and coastal management. In this research I used remote sensing, field observations and numerical modelling in the Plum Island Sound in Massachusetts, USA, to explore mechanisms controlling sediment dynamics and their feedbacks with SLR. The analysis of remote-sensed suspended sediment concentrations (SSC) reveals that a 5-year record (2013-2018) is sufficient to capture a representative range of meteorological and tidal conditions required to determine the main drivers of SSC dynamics in hydrodynamically-complex and small-scale coastal bays. The interplay between river and tidal flows dominated SSC dynamics in this estuary, whereas wind-driven resuspension had a more moderate effect. The SSC was higher during spring because of increased river discharge due to snowmelt. Tidal asymmetry also enhanced sediment resuspension during flood tides, possibly favoring deposition on marsh platforms. Together, water level, water-level rate of change, river discharge and wind speed were able to explain > 60% of the variability in the main channel SSC, thereby facilitating future prediction of SSC from these readily available variables. To determine the fate of cohesive sediments and spatial variations of trapping capacity in the system, a high-resolution (20 m) numerical model coupled to a vegetation module was developed. The results highlight the importance of the timing between sediment inputs and tidal phase and show that sediment discharged from tidal rivers deposit within the rivers themselves or in adjacent marshes. Most sediment is deposited in shallow tidal flats and channels and is unable to penetrate farther inside the marshes because of the limited water depths and velocities on the marsh platform. Trapping capacity of sediment in different intertidal subdomains decreases logarithmically with the ratio between advection length and the typical length of channels and tidal flats. Moreover, sediment deposition on the marsh decreases exponentially with distance from the channels and marsh edge. This decay rate is a function of settling velocity and the maximum value of water depth and velocity on the marsh platform. Bed sediment compositions were generated to further explore feedbacks between SLR, sediment dynamics and morphological changes. The results show SLR increases tidal prism and inundation depth, facilitating sediment deposition on the marsh platform. At the same time, SLR enhances ebb-dominated currents and increases sediment resuspension, reducing the sediment-trapping capacity of tidal flats and bays, leading to a negative sediment budget for the entire system. This bimodal distribution of sediment budget trajectories will have a profound impact on the morphology of coastal bays, increasing the difference in elevation between salt marshes and tidal flats and potentially affecting intertidal ecosystems. The results also clearly indicate that landforms lower with respect to the tidal frame are more affected by SLR than salt marshes. Therefore, Salt marshes, shallow bays, tidal flats, and barrier islands are inherently and physically connected systems, and evaluating the effect of SLR on salt marshes should involve all these units.

No description available.

Thesis advisor: Gail Kineke / Thesis advisor: Noah Snyder / Salt marshes are some of the world’s richest ecosystems and provide a plethora of benefits to coastlines and bays in terms of storm protection and chemistry. To ensure salt marsh survival under increasing rates of sea level rise, management practices have been trending towards natural sustainability measures to increase marsh resilience. To benefit these efforts, it is necessary to understand how natural salt marshes respond to environmental change in terms of sediment deposition and evolution of vegetation and open water. This study uses aerial image digitization to understand how Nauset Marsh in Cape Cod MA, a protected salt marsh on Cape Cod National Seashore, has responded to sea level rise and half a century of inlet migration. Digitized images from 1974-2019 were used to track changes to vegetation extent and open water features during study periods of different inlet migration stages. Observed changes were used to ascertain trends of marsh loss or adaptation based on previous research on ponding cycles and vegetation extent. Results indicate that Nauset Marsh has been relatively stable over the last half century, with the most significant change observed in Vegetated Marsh loss of 6.71% ± 3.19 primarily due to edge erosion near the present-day inlet. Despite net feature stability, significant differences in feature evolution trends were observed during different stages of inlet migration. Most notably, inlet breaching and migration correlated with dynamic feature changes throughout the marsh, while the static inlet period correlated with expansion of open water features near the inlet location. The evolution of Nauset Marsh suggests that inlet migration improves marsh resilience through periodic increases in sediment deposition in a natural salt marsh with sufficient sediment supply. / Thesis (MS) — Boston College, 2023. / Submitted to: Boston College. Graduate School of Arts and Sciences. / Discipline: Earth and Environmental Sciences.

Appreciation of the ecological and economic values associated with healthy salt marshes has led to a recent rise in the number of marshes that are being targeted for restoration by dike removal. The success of restoration is often measured by the return of marsh plants, though this overlooks a key component of salt marshes, that of the invertebrate community within marsh sediments. To evaluate the short-term recovery of these invertebrates, sediment cores were collected across an elevational gradient in a recent dike removal marsh, one and two years post removal, and a nearby reference marsh. Abundance, richness and diversity as well as morphospecies community composition were compared across treatment groups (Reference, Removal) and elevation zone (High Marsh, Low Marsh). Morphospecies richness, abundance and diversity were significantly higher in Low Marsh samples than in High Marsh samples, though no statistically significant differences were found across treatments of the same elevation (e.g., Reference Low Marsh versus Removal Low Marsh). Pair-wise ANOSIM results found significant differences between community compositions across treatments, specifically Reference Low Marsh and Removal Low Marsh. The marsh edge, the lowest point of vascular plant growth before transitioning to tide flats, is considered a high stress environment for emergent vegetation. Plant establishment and survival in this low elevation zone is limited by the tolerance to inundation duration and frequency and anoxic sediments. Bioturbation and burrowing by macroinvertebrates increases the surface area exposed to surface water for gas exchange, increasing the depth of the redox potential discontinuity layer. Crabs that make stable, maintained burrows have been shown to increase oxygen penetration into sediment, improving plant productivity. Such crabs are not found in salt marshes of the Pacific Northwest of North America. However, other burrowing invertebrates may have a positive impact on plant health in these areas by reducing abiotic stress due to anoxic sediments, thereby allowing plants to establish and survive lower in the intertidal zone. To assess this potential relationship, study plots of Distichlis spicata were selected at equivalent elevations at the lowest point of plant establishment at the marsh edge. Focal plant rhizomes were severed from upland ramets and assigned an invertebrate abundance treatment based on a visual burrow count surrounding each plant (9 cm diameter). Focal plants were visited monthly from July to September 2016, plant health variables of chlorophyll content and chlorophyll fluorescence (photosynthetic efficiency), and sediment ORP readings were collected. Plant survivorship was significantly higher in plots with invertebrates, 96% of plants in 'With Invertebrate' plots and 50% of plants in 'No Invertebrates' plots survived the duration of the study. Plant health (chlorophyll content and chlorophyll fluorescence) generally increased with increased invertebrate presence though, not statistically significant. There may be potential for improved plant productivity and resilience to plants at the marsh edge due to invertebrate burrowing activity. This benefit could help mitigate projected losses in plant productivity due to sea level rise, though more research is needed to investigate the mechanism by which these invertebrates confer a health benefit to plants at the marsh edge.

Herbivores and pathogens pose a consistent threat to plant productivity. In response, plants invest in structural and/or chemical defenses that minimize damage caused by these biotic stressors. In salt marshes along the Atlantic coast of the United States, a facultative mutualism between snails (Littoraria irrorata) and multiple species of fungi exert intense top-down control of the foundation grass species Spartina alterniflora. Since exposure to herbivores and pathogens are tightly coupled in this system, I investigated whether S. alterniflora utilizes chemical and/or structural defenses to deter both snails and fungi, and examined how plant defenses varied among S. alterniflora individuals and populations. I also assessed how other marsh plants prevent snails from establishing farms, and considered whether interspecific variation in plant chemical defenses influences marsh community structure. Initial experiments revealed that S. alterniflora chemical defenses inhibited L. irrorata and two fungi that snails commonly farm. A caging experiment determined that production of chemical defenses could not be induced in the presence of snails and fungi, nor relaxed in their absence. Through separations chemistry guided by ecological assays, I isolated two distinct classes of chemical defenses from short form S. alterniflora, one of which inhibited fungal growth and the other decreased plant palatability. In a community context, the chemical defenses produced by S. alterniflora were relatively weak compared to those of four other salt marsh plant species, which produced compounds that completely inhibited L. irrorata grazing and strongly hindered fungal growth in lab assays. Nutritional and structural differences among marsh plants did not influence feeding preferences, suggesting that plant secondary chemistry was the primary driver for food selection by snails. It appears that S. alterniflora produces weak chemical defenses that slow down or limit fungal growth and snail herbivory, and may compensate for tissue losses by producing new growth. In contrast, less abundant marsh plants express chemical defenses that completely inhibit fungal farming and deter snail grazing, but doing so may come at a cost to growth or competitive ability. As marsh dieback continues with rising herbivore densities and compounding abiotic stressors, the ecosystem services that salt marshes provide may be lost. Therefore, understanding how and under what conditions salt marsh plants resist losses to herbivores and pathogens will help predict which marsh communities are most likely to be threatened in the future. Initial experiments revealed that S. alterniflora chemical defenses inhibited L. irrorata and two fungi that snails commonly farm. A caging experiment determined that production of chemical defenses could not be induced in the presence of snails and fungi, nor relaxed in their absence. Through separations chemistry guided by ecological assays, I isolated two distinct classes of chemical defenses from short form S. alterniflora, one of which inhibited fungal growth and the other decreased plant palatability. In a community context, the chemical defenses produced by S. alterniflora were relatively weak compared to those of four other salt marsh plant species, which produced compounds that completely inhibited L. irrorata grazing and strongly hindered fungal growth in lab assays. Nutritional and structural differences among marsh plants did not influence feeding preferences, suggesting that differences in plant chemistry were the primary driver for food selection by snails. It appears that S. alterniflora produces weak chemical defenses that slow down or limit fungal growth and snail herbivory, and may compensate for tissue losses by producing new growth. In contrast, less abundant marsh plants express chemical defenses that completely inhibit fungal farming and deter snail grazing, but doing so may come at a cost to growth or competitive ability against S. alterniflora. As marsh dieback continues with rising herbivore densities and compounding abiotic stressors, the ecosystem services that salt marshes provide may be lost. Therefore, understanding how and under what conditions salt marsh plants resist losses to herbivores and pathogens will help predict which marsh communities are most likely to be threatened in the future.

The ability of prey to detect and respond appropriately to predator risk is important to overall prey fitness. Many aquatic organisms assess risk through the use of chemical cues that can change with predator diet. Two variable characteristics of diet are: 1. prey type and 2. prey mass. To assess the effect of these two characteristics on the assessment of risk by the mud crab Panopeus herbstii, I exposed mud crabs to the urine of the blue crab Callinectes sapidus fed one of 5 diet treatments: 10g of oyster shell free wet mass, 5g of oyster shell free wet mass, 10g crushed mud crabs, 5g crushed mud crabs, and a mix of 5g of oyster shell free wet mass and 5g crushed mud crab. Effects on P. herbstii foraging were tested in a previously developed bioassay by measuring shrimp consumption over a 4 hour period. I hypothesized that P. herbstii would have a larger magnitude response to urine from C. sapidus fed a diet of crushed mud crabs than to urine from C. sapidus fed a diet of oysters. I further hypothesized that P. herbstii would have a larger magnitude response to urine from C. sapidus fed a high mass diet relative to a lower mass diet. Contrary to expectations there was no observed effect of urine on P. herbstii foraging in any of the treatments. Results suggest that bioassay protocol may be unreliable suggesting further replication to determine the difference between this study and previous results. Future studies examining how P. herbstii varies with urine concentration will aid in understanding the ecological scale of this predator cue system. Determining the role of other potential cue sources will improve the predictive abilities of these studies.

We quantified U removal and investigated the efficacy of uranium as a quantitative tracer of groundwater discharge in a headwater salt marsh of the Okatee River, Bluffton, SC. Determining the magnitude of U removal is important for advancing U as a tracer of paleo-oceanic conditions. Since salt marsh groundwater is typically enriched in nutrients and other biologically and chemically reactive species, quantifying groundwater discharge from marshes is critical for understanding the ability of salt marshes to modify the chemistry of important species in surface waters. We hypothesized that water-column U(VI) was removed by tidally-induced advection of surface water into permeable, anoxic salt marsh sediments, a process resulting in bacterially-mediated precipitation of insoluble U(IV)O2 and/or sorption of uranium to iron-oxides at the oxic/anoxic sediment interface. Furthermore, we suggested that hydraulic pressure gradients established by marsh-surface tidal inundation and seasonally-variable rainfall promote the discharge of salt-marsh-processed, uranium-depleted groundwater to tidal creeks, producing the surface-water U-removal signal. Groundwater and surface water data revealed non-conservative uranium behavior. We documented extensive uranium removal from shallow marsh groundwater and seasonally variable uranium removal from surface waters. These observations allowed for the calculation of seasonally-dependent salt marsh uranium removal rates. On a yearly basis, our removal rate (58 to 104 mol m-2 year-1) reemphasized the importance of anoxic coastal environments for U removal. High uranium removal, high barium concentration water observed seeping from creek banks at low tide supported our hypothesis that groundwater discharge must contribute to uranium removal documented in tidal surface waters. Average site groundwater provided an analytically reasonable endmember for explaining uranium depletion in surface water. Therefore, we used three endmember mixing models for estimating the fraction of surface water with presumed a groundwater signature. Our discharge estimates of 8 to 37 L m-2 day-1 agreed closely with previously published salt marsh values. Seasonality in discharge rates can be rationalized with appeal to seasonal patterns in observed rainfall, tidal forcing, and marsh surface bioturbation. Although more work is needed, the results of this portion of the study suggest that U may be an effective quantitative tracer of groundwater discharge from salt marshes.